Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
A ailable online 5 Oc obe 2023
0031-0182/© 2023 Else ie B.V. All igh s ese ed.
Laguna Seca sedimen s e eal en i onmen al and clima e change du ing
he la es Pleis ocene and Holocene in Sie a Ne ada, sou he n
Ibe ian Peninsula
Gonzalo Jim´
enez-Mo eno
a
,
*
, Alejand o L´
opez-A il´
es
a
, An onio Ga cía-Alix
a
,
b
,
Ma ía J. Ramos-Rom´
an
c
, Jon Camue a
b
, Jose Manuel Mesa-Fe n´
andez
a
,
F ancisco J. Jim´
enez-Espejo
b
, Cha o L´
opez-Blanco
a
, Jos´
e S. Ca i´
on
d
, R. Sco Ande son
e
a
Depa amen o de Es a ig a ía y Paleon ología, Uni e sidad de G anada, Spain
b
Ins i u o Andaluz de Ciencias de la Tie a (IACT), CSIC-UGR, A milla, Spain
c
Facul ad de Educaci´
on, Uni e sidad del A l´
an ico Medio, Mad id, Spain
d
Depa men o Plan Biology, Facul y o Biology, Uni e si y o Mu cia, Spain
e
School o Ea h and Sus ainabili y, No he n A izona Uni e si y, USA
ARTICLE INFO
Edi o : M Ellio
Keywo ds:
Vege a ion
Lake le el
Clima e
La e Pleis ocene
Holocene
Wes e n Medi e anean
ABSTRACT
Sedimen a ion in mos glacial lakes and we lands in he Sie a Ne ada (sou he n Ibe ian Peninsula) began a e
he las deglacia ion and since he Younge D yas (YD)-Ea ly Holocene (EH) ansi ion. The e o e, un il now,
s udies on olde sedimen a y eco ds we e lacking in his alpine a ea, which limi s he paleoen i onmen al and
paleoclima ic in o ma ion o he Holocene. In his s udy, we s udied palynomo phs om he alpine eco d om
Laguna Seca (LS), he longes and oldes (~18,000 cal y BP =18 ky ) sedimen a y eco d in he Sie a Ne ada o
in es iga e he esponse o o es s and lake en i onmen s in he wes e n Medi e anean a ea o clima e changes
and human impac du ing he la es Pleis ocene and Holocene. The deepes lake condi ions occu ed du ing he
las deglacia ion, indica ed by he occu ence o Pedias um algae, which showed highes abundances du ing he
Hein ich S adial 1 (HS1) and Bølling-Alle ød (B-A) ansi ion. Xe ophy e he bs such as A emisia, Ephed a, and
Ama an haceae we e highes du ing he la e B-A and YD indica ing egional a idi y. Poaceae (g asses) we e
maxima in he B-A and EH, p obably indica ing expansion in he ba en a eas a e deglacia ion. Maximum in
empe a u e and humidi y du ing he EH and cooling and a idi ica ion in he Middle (MH) and La e Holocene
(LH) a e indica ed by he changes in he abundance o deciduous Que cus and Pinus o es species. Bo yococcus
algae inc eased du ing he Ea ly Holocene, while he es o he algae almos anished, which could indica e ha
he lake became e y p oduc i e bu shallowe un il 8.2 ky . The lake le el lowe ed and became seasonal in he
Middle-La e Holocene ansi ion, coinciding wi h he egional clima e a idi ica ion. Mic ocha coal analysis done
on he palynological p epa a ions ag ees wi h he ege a ion changes, showing maxima in he EH and MH,
ela ed wi h he maximum in egional o es occu ence, and a dec ease in he LH when he Medi e anean
ege a ion, and hus uel a ailabili y, diminished. This eco d shows e idence o an h opogenic impac in he las
cen u ies by cul i a ion, e o es a ion, ca le g azing, enhanced e osion and eu ophica ion.
1. In oduc ion
The Sie a Ne ada is one o he highes ele a ion and sou he nmos -
loca ed moun ain ange in sou he n Eu ope (Fig. 1). I lies wi hin a
pa icula biogeog aphic egion (Zamo a and Oli a, 2022), enowned as
a biodi e si y supe ho spo (A oyo e al., 2022). High-ele a ion
moun ain en i onmen s can be pa icula ly sensi i e o ecen clima e
wa ming and inc ease in d ough condi ions (P´
ascoa e al., 2017), which
is causing al i udinal displacemen s o plan and animal species, and loss
o biodi e si y in he Sie a Ne ada (Lo i e e al., 2022). Wi h p o-
jec ions o annual mean wa ming on land o 0.9–5.6 ◦C a he end o he
cen u y in he Medi e anean a ea (Ali e al., 2022), wha will be he
en i onmen al consequences o such a apid clima e change in moun-
ains such as he Sie a Ne ada?
* Co esponding au ho .
E-mail add ess: [email p o ec ed] (G. Jim´
enez-Mo eno).
Con en s lis s a ailable a ScienceDi ec
Palaeogeog aphy, Palaeoclima ology, Palaeoecology
jou nal homepage: www.else ie .com/loca e/palaeo
h ps://doi.o g/10.1016/j.palaeo.2023.111834
Recei ed 20 Ap il 2023; Recei ed in e ised o m 11 July 2023; Accep ed 2 Oc obe 2023
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
2
Fig. 1. (A) Loca ion o Sie a Ne ada ( ec angle) wi hin he Ibe ian Peninsula, wes e n Medi e anean egion. The MD95–2043 and ALB-2 ma ine si es discussed in
he ex a e also shown. Sa elli e pho o om Google Ea h. (B) Loca ion o Laguna Seca (LS) wi hin he Sie a Ne ada. The Padul and Sie a Ne ada si es discussed in
he ex a e also shown (1: Bo eguil de la Vi gen; 2: Laguna de Río Seco; 3: Laguna de la Mosca; 4: Laguna Honde a). Sa elli e pho o om Google Ea h. (C) Wind
di ec ion in pe cen age om Janua y 2000 o Decembe 2019 o G anada, Spain (da a ob ained om h ps://www.woeu ope.eu) (D) LS a ea and opog aphy. No e
he eeline immedia ely sou h o he lake a ~2300 m. Sa elli e pho o om Google Ea h. (E) LS du ing he co ing o LS-01 in Sep embe 2014. (F) LS con aining
wa e in ea ly June 2018. (G) Map o he ege a ion bel s in he Sie a Ne ada (Modi ied om REDIAM. Map o he ege a ion se ies o Andalucía: h p://labo a o i
o ediam.cica.es/Viso Gene ico/? ipo=WMS). Figu e modi ied om Ramos-Rom´
an e al. (2018a) and L´
opez-A il´
es e al. (2022).
G. Jim´
enez-Mo eno e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
3
Paleoecological and paleoclima ic s udies a e necessa y o p edic
he u u e, imp o ing ou unde s anding o he ela ionship be ween
clima e and moun ain ecosys ems in he pas , especially du ing wa me
and d ie pe iods han a p esen and du ing signi ican clima ic luc-
ua ions such as he glacial-in e glacial ansi ions. In his espec , he e
a e mo e han 50 small glacial-o igin alpine lakes and bogs in he Sie a
Ne ada ha accumula ed sedimen s p ese ing a high-quali y signal o
pas na u al and an h opogenic en i onmen al and clima e change
(Ande son e al., 2011; Jim´
enez-Mo eno and Ande son, 2012; Ga cía-
Alix e al., 2012, 2013, 2017, 2018, 2020; Jim´
enez-Espejo e al., 2014;
Jim´
enez-Mo eno e al., 2013, 2020; Ramos-Rom´
an e al., 2016, 2018a,
2018b, 2019; Mesa-Fe n´
andez e al., 2018; Manzano e al., 2019; Toney
e al., 2020; L´
opez-A il´
es e al., 2021; Alba-S´
anchez e al., 2021). P e-
ious s udies show ha in he glacial lakes and we lands om he Sie a
Ne ada sedimen a ion s a ed a e deglacia ion and he ea e since he
Ea ly Holocene (see syn hesis in Jim´
enez-Mo eno e al., 2022). The e-
o e, olde sedimen a y eco ds we e ye o be ound in his alpine a ea
o widen ou knowledge abou en i onmen al change u he in ime
and since he las deglacia ion.
In 2014 h ee sedimen a y co es, LS-01, LS-02 and LS-03, we e ob-
ained om LS (L´
opez-A il´
es e al., 2022; Fig. 1), s ill in he alpine a ea
bu a lowe ele a ion han he p e ious s udies. The age model o he
14 m-long sedimen a y sequence, based on nine Accele a o Mass
Spec ome y (AMS) adioca bon and ou op ically s imula ed lumi-
nescence (OSL) da es, shows a con inuous eco d o he las ~18 ky and
since he Las Glacial Maximum (LGM) (L´
opez-A il´
es e al., 2022;
Fig. 2). In his s udy, we analyzed palynomo phs and cha coal abun-
dance om he LS-01 alpine eco d, which ep esen s so a , he longes
and oldes sedimen a y eco d in Sie a Ne ada, sou he n Spain. Ou
goal is o documen ege a ional de elopmen s h ough palynological
and cha coal da a and join hose o p e iously epo ed ch onological,
sedimen ological and geochemical in o ma ion by L´
opez-A il´
es e al.
(2022). Fu he mo e, he compa ison o he LS-01 eco d wi h de ailed
paleoen i onmen al and paleoclima ic eco ds om he wes e n Medi-
e anean allows us o ob ain in o ma ion on po en ial clima ic and
an h opogenic igge s o pas en i onmen al change in he s udy a ea.
2. S udy si e
LS (37◦05
′
53”N, 2◦58
′
05”W, 2259 m abo e sea le el) is a small
endo heic alpine lake basin loca ed in he eas e n sec o o he Sie a
Ne ada acing sou h igh below he idge line (Fig. 1). LS o med om
he combina ion o ec onics, including a o a ional aul , and subse-
quen glacial and pe iglacial e osion, indica ed by he occu ence o
mo aines in he LS alley (Sim´
on e al., 2000). A p esen , LS ge s
co e ed wi h ice and snow in win e and becomes inunda ed in sp ing
and ea ly summe , when he lake le el can be as high as 50 cm (pe sonal
obse a ions be ween 2014 and 2023). LS is seasonally d y du ing
summe and ea ly all (Fig. 1). The lake wa e supply comes om
snowmel om he d ainage a ea a ound he lake, and occasional uno
Fig. 2. LS-01 co e li hology, magne ic suscep ibili y (MS in SI uni s x 10
−4
) and bayesian age model (wi h es ima ed sedimen a y a es in mm/y ). The sec ion o he
co e s udied he e o he opmos 4.7 m is shown in ed in he li hological plo . On he igh is a plo wi h down-co e unce ain y anges plo ed on age. Fo mo e
de ails abou he li hology and age model see L´
opez-A il´
es e al. (2022). (Fo in e p e a ion o he e e ences o colou in his igu e legend, he eade is e e ed o
he web e sion o his a icle.)
G. Jim´
enez-Mo eno e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
4
e en s. The LS basin bed ock is composed o low-deg ee me amo phic
mica schis s o he Ne ado Fil´
ab ide complex (Ma ín Ma ín e al.,
2010).
The clima e in he s udy a ea is ypically Medi e anean, wi h mild
win e s and ho and d y summe s. Clima ic da a o LS a e no di ec ly
a ailable, bu a wea he s a ion loca ed ~3.5 km om he s udy si e a
2200 m ele a ion shows a mean annual p ecipi a ion o 581 mm and a
mean annual empe a u e o 6.3 ±0.5 ◦C. The d ies mon h is July wi h
2 mm and he mos humid mon h is Ma ch wi h 114 mm o ain all.
Feb ua y is he coldes mon h (−1.2 ±1.7 ◦C) and July he wa mes one
(16.8 ±1.2 ◦C) (O ganismo Au ´
onomo Pa ques Nacionales, 2021).
The p esen -day ege a ion in he Sie a Ne ada is la gely con olled
by empe a u e and p ecipi a ion/humidi y g adien s and hus depends
on he ele a ion (Valle, 2003; Fig. 1G). Abo e ~2800 m asl, clima e is
cha ac e ized by e y cold win e s, sho g owing season, high sola
adia ion and snow all, and s ong winds. In his con ex , soils a e
meage and a und a-like sh ubby g assland de elops. This c yo -
omedi e anean ege a ion bel is cha ac e ized by species o Poaceae,
As e aceae, B assicaceae and Plan aginaceae be ween o he he bs,
including se e al endemic plan s (E ige on igidus, Saxi aga ne adensis
and Viola c assiuscula) (Valle, 2003). The unde lying o omedi e anean
ege a ion bel is loca ed be ween ~1900–2800 m asl, and is mos ly
cha ac e ized by coni e s, such as Pinus syl es is, P. nig a, Junipe us
hemisphae ica, J. sabina and J. communis subsp. nana, as well as woody
species o Fabaceae, Cis aceae and B assicaceae. The sup a-
medi e anean bel , be ween ~1400–1900 m asl, is cha ac e ized by
Que cus py enaica, Q. aginea, Q. o undi olia, Ace opalus subsp.
g ana ense and F axinus angus i olia among o he ees/sh ubs and some
he bs such as A emisia glu inosa. The mesomedi e anean ege a ion
bel , be ween ~600–1400 m asl, is mainly domina ed by Que cus
o undi olia wi h a cha ac e is ic Re ama sphae oca pa unde s o ey
(Valle, 2003). Que cus cocci e a and Pis acia len icus a e he main axa in
he lowes ele a ion he momedi e anean ege a ion bel (~0–600 m
asl). LS is loca ed a he uppe ele a ional limi o he o omedi e anean
ege a ion bel and eeline in he s udy a ea, loca ed a ~2300 m asl,
mos ly composed o P. syl es is (Fig. 1D). Howe e , he s udy a ea has
been modi ied by high an h opogenic ac i i y and was e o es ed wi h
P. syl es is in he mid-20 h cen u y (A ias Abell´
an, 1981). The e o e, i
is di icul o place he na u al ele a ion o eeline in Sie a Ne ada,
which seems o ha e been a i icially lowe ed down se e al hund ed
me e s (Lo i e e al., 2022).
3. Ma e ials and me hods
The 14.1 m-long LS-01 sedimen a y eco d was eco e ed om he
depocen e o he LS basin in Sep embe 2014 using a Rola ec RL-48-L
co ing machine om he Scien i ic Ins umen a ion Cen e o he Uni-
e si y o G anada (CIC-UGR) (Fig. 1). The co es we e anspo ed o he
UGR, s o ed in a e ige a ed oom a 4 ◦C and e en ually we e spli and
imaged in he labo a o y a UGR and CIC, espec i ely. Li hologic
desc ip ion and sedimen ological analysis we e p e iously ca ied ou
by L´
opez-A il´
es e al. (2022) (Fig. 2). To de elop an age model o he
LS-01 sedimen co e, we used adioca bon analyses om he opmos
o ganic- ich pa o he eco d and OSL analyses o he de i al- ich
Table 1
Age da a o LS-01. Radiome ic ages we e calib a ed using In Cal20.14c cu e (Reime e al., 2020) wi h
Calib 8.2 (h p://calib.o g/calib/). OSL da es a e ma ked wi h as e isk. In ed colou a e he adioca bon
da es no used in he age model. Ac onyms: TBS; p e- ea ed bulk sedimen wi h HCl and HF, BS; bulk
sedimen .
G. Jim´
enez-Mo eno e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
5
bo om (L´
opez-A il´
es e al., 2022; Fig. 2). P e ious analyses on he LS-01
co e included magne ic suscep ibili y (MS), o al o ganic ca bon and C/
N (L´
opez-A il´
es e al., 2022; Fig. 2).
The opmos ~4.7 m pa o he LS-01 sedimen a y eco d, cha ac-
e ized by ine-g ained sedimen s, was sampled o palynological anal-
ysis e e y 5 cm. This sec ion o he co e con ains a sedimen a y eco d o
he las ~15.8 ky based on he age model desc ibed in L´
opez-A il´
es
e al. (2022) (Table 1; Fig. 2). A o al o 94 samples o wo cubic cen-
ime e s we e collec ed. The pollen ex ac ion me hod ollowed a
modi ied Faeg i and I e sen (1989) me hod. One able wi h a known
numbe o Lycopodium spo es o calcula ion o pollen concen a ion was
added o he samples. Sedimen samples we e p ocessed wi h HCl (35%)
and HF (45%) o emo e he ca bona es and silica es, espec i ely, om
he sedimen and concen a e he o ganic ma e . The samples we e
sie ed a 10
μ
m o emo e pa icles smalle han he pollen g ains. The
esidual ma e ial, oge he wi h glyce in, was moun ed on mic oscope
slides. Coun ing, and iden i ica ion, o palynomo phs was ca ied ou
wi h a Leica DM1000 LED ansmi ed ligh mic oscope a 400×mag-
ni ica ions. Pollen a lases, such as Beug (1961) and pollen e e ence
ma e ial o plan s ha occu in sou he n Spain we e used o ce ain
iden i ica ions. Pinus pollen was di ided in o wo ca ego ies: Pinus
inde e mina e and P. syl es is ype (mos -likely including P. syl es is
and/o P. nig a). P. syl es is ype pollen was di e en ia ed om he es
o Pinus inde e mina e pollen by i s small size (gene ally wi h a o al
g ain size smalle han 85
μ
m) (Desp a e al., 2015). Pollen pe cen ages
in each analyzed sample ha e been calcula ed wi h espec o he o al
sum o e es ial pollen. The mos ep esen a i e pollen ypes wi h
occu ences highe han 1% a e shown in Fig. 3. The pe cen ages o
non-pollen palynomo phs (NPP), including algae such as Bo yococcus,
Pedias um, Deba ya, Spi ogy a and Zygnema, dino lagella e cys s (dino-
cys s), Isoe es mic ospo es, oocy es o he aqua ic la wo m Neo-
habdocoela, myco hizal chlamydospo es o Glomus and spo es o he
li e wo Riccia ha e been calcula ed wi h espec o he o al sum o
e es ial pollen plus NPP and a e shown in Fig. 4. The pollen clima e
index (PCI), used in p e ious s udies o disc imina e be ween cold/a id
and wa m/humid clima es (e.g., Joannin e al., 2011; Camue a e al.,
2019, 2021), and calcula ed wi h he a io o meso he mic (Que cus,
Olea, Be ula and Co ylus) / s eppic axa (A emisia, Lygeum, Ephed a and
Ama an haceae), was also calcula ed he e (Fig. 5). Clus e analyses o
he pollen and NPP da a we e pe o med using he p og am CONISS
(G imm, 1987), wi h he aim o g ouping samples wi h simila pollen
assemblages in o pollen zones (Figs. 3 and 4).
Cha coal pa icles bigge han 30
μ
m we e coun ed in he palyno-
logical slides. Cha coal concen a ion (cha coal pa icles/cm
3
) was
calcula ed using he Lycopodium ace concen a ion (Figs. 3 and 5) in
his way: #cha coal pa icles/#Lycopodium ace s *9666 (being 9666
he numbe o Lycopodium ace s in 1 able ).
4. Resul s
The LS-01 pollen eco d is made up o 42 samples ha p o ided
su icien pollen o s a is ical analyses. Fi y- wo samples we e ba en
in pollen. The pollen eco d is gene ally cha ac e ized by he ele ance
o he o es componen , which is domina ed by Pinus species, mos
likely including P. syl es is ( ha g ows in he a ea a p esen ) o P. nig a.
To a lesse ex en , he pollen spec a show e e g een and deciduous
Que cus, Junipe us, Olea, Be ula and Co ylus. Rega ding he he baceous
and sh ubby axa, pollen assemblages a e domina ed by Poaceae, A e-
misia, As e aceae As e oideae, As e aceae Cicho ioideae, Ama -
an haceae, Ca yophyllaceae, Plan ago, and in smalle p opo ions,
E icaceae, Ephed a, Lygeum, Rosaceae, Onag aceae, Ge aniaceae and
Cis aceae. Cype aceae and Ranunculaceae a e also ela i ely abundan
and ep esen he aqua ic plan s (Fig. 3). Wi h espec o he NPP,
Pedias um, Bo yococcus, Spi ogy a, Zygnema and Deba ya, a e abundan .
Isoe es, Riccia, Glomus, Neo habdocoela and dinocys s also occu (Fig. 4).
Cha coal concen a ion a ies conside ably h ough ime (Fig. 3).
The pollen and NPP equencies luc ua e conside ably h ough ime
and can be subdi ided in o se e al en i onmen al and clima ically
dis inc pe iods (zones) ha oughly ag ee wi h he subdi ision o he
la es Pleis ocene (HS1, B-A and YD; Rasmussen e al., 2006) and Ho-
locene Se ies/Epoch (Ea ly, Middle and La e Holocene; Walke e al.,
2018; Figs. 3 and 4). Below, we desc ibe he LS pollen and NPP da a o
each o he iden i ied zones.
4.1. Pollen and NPP zone 1 (be ween ~ 15.8–14.2 ky ; 470–405 cm
dep h)
This palynological zone migh oughly co esponds in iming wi h
he HS1 and B-A ansi ion. P. syl es is / nig a, shows maxima du ing
his zone wi h pe cen ages a e aging ~95%. As e aceae As e oideae,
A emisia and Ama an haceae inc eased sligh ly a he end o his zone
(Fig. 3). Cha coal shows ela i ely low concen a ions. The aqua ic alga
Pedias um is mos abundan in his zone p esen ing wo peaks. The alga
Bo yococcus peaks du ing he Pedias um minimum. Zygnema aceae
zygospo es (Deba ya, Spi ogy a and Zygnema) a e abundan . Neo-
habdocoela occu s wi h maximum equencies in his zone (Fig. 4).
4.2. Pollen and NPP zone 2 (be ween ~ 14.2–12.7 ky ; 405–370 cm
dep h)
This palynological zone app oxima ely co esponds o he B-A
(Figs. 3 and 4). Pinus dec eases conside ably, ini ially eaching ~40%.
Fig. 3. De ailed pollen and cha coal diag am om he LS-01 co e showing pe cen ages o selec ed axa (highe han 1%). In g een, yellow and blue a e he ees,
he bs and we land angiospe m axa, espec i ely. The pollen zona ion on he igh was made using clus e analysis p o ided by CONISS (G imm, 1987). Ligh
shadings indica e a 5×exagge a ion scale. HS1, B-A, YD, EH, MH and LH s and o Hein ich S adial-1, Bolling-Alle od, Younge D yas, Ea ly Holocene, Middle
Holocene and La e Holocene. (Fo in e p e a ion o he e e ences o colou in his igu e legend, he eade is e e ed o he web e sion o his a icle.)
G. Jim´
enez-Mo eno e al.

Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
6
Subsequen ly, i shows wo oscilla ions, wi h peaks ~90% and a mini-
mum o ~40%. Junipe us inc eased a he beginning o his zone. He -
baceous axa such as Poaceae, A emisia, Ephed a and Ama an haceae
inc ease. A emisia shows maximum alues a he end o his zone.
Cha coal shows highe abundance. Pedias um and Bo yococcus,
oge he wi h he Zygnema aceae and Neo habdocoela, dec ease
conside ably, whe eas dinocys s show maximum alues.
4.3. Pollen and NPP zone 3 (be ween ~ 12.7–11.8 ky ; 370–350 cm
dep h)
This zone is co ela ed wi h he YD (Figs. 3 and 4). Pinus shows
ma ked a iabili y, wi h wo maxima and wo minima wi h alues be-
ween 95 and 20%. He bs such as A emisia, Plan ago and Ama -
an haceae peak in his zone, wi h he la es showing maximum alues.
The aqua ic Ranunculaceae inc ease, showing a peak a his ime.
Cha coal pa icles dec ease and show minimum alues. Wi h espec o
he NPP, he mic oalgae Bo yococcus and Zygnema aceae (including
Deba ya, Spi ogy a and Zygnema), inc ease. Neo habdocoela is absen
while he dinocys s decline.
4.4. Pollen and NPP zone 4 (be ween ~ 11.8–8.4 ky ; 350–290 cm
dep h)
This zone app oxima ely co esponds o he EH (Figs. 3 and 4). Pinus
is luc ua ing wi h a dec easing end, eaching wo minima o ~5% a
10.5 and 8.4 ky . Junipe us and bo h e e g een and deciduous Que cus
inc ease in his zone. Deciduous Que cus shows he eco d’s maximum a
~10.6 ky . E icaceae, Be ula and Co ylus also inc ease, wi h he la e
showing an inc easing end and maximum alues a he end o he EH.
He bs such as Poaceae, Ca yophyllaceae, Onag aceae and Ge aniaceae
inc ease, whe eas A emisia dec ease, showing minimum alues o he
eco d be ween ~10.5–8.4 ky . The aqua ic Cype aceae show maximum
occu ences du ing he EH, peaking oge he wi h Ranunculaceae a
~11 ky . Cha coal concen a ion inc ease conside ably, showing max-
ima be ween 11 and 10.2 and 9.4–8.4 ky . Bo yococcus inc ease
signi ican ly du ing he EH whe eas o he algae such as Zygnema aceae
(including Deba ya, Spi ogy a and Zygnema) dec ease. Neo habdocoela
only occu a he beginning o his zone and Isoe es occu s wi h a peak a
he end o he EH.
4.5. Pollen and NPP zone 5 (be ween ~ 8.4–5.6 ky ; 290–170 cm dep h)
This zone co esponds o mos o he MH (Figs. 3 and 4) and is
cha ac e ized by he minimum alues o Pinus, a e aging 20%. O he
o es species such as e e g een and deciduous Que cus, Junipe us and
Olea a e ela i ely abundan du ing his pe iod. Wi h espec o he
he bs, A emisia and Ca yophyllaceae inc ease in his zone. The aqua ic
Ranunculaceae shows maximum alues o he eco d whe eas Cype -
aceae dec ease. Cha coal concen a ion is cha ac e ized by a peak a he
beginning o he MH a ~8.0 ky bu declines du ing he es o he MH.
The NPP a e mos ly ep esen ed by Bo yococcus, which shows
maximum occu ences be ween 8.4 and 7.8 ky , dec easing conside ably
du ing he es o he MH.
4.6. Pollen and NPP zone 6 (be ween ~ 5.6 ky -p esen ; 170–0 cm
dep h)
This palynological zone co esponds o he la es MH and La e Ho-
locene (LH; Figs. 3 and 4). I is cha ac e ized by he inc ease in Pinus,
depic ing a peak a ~5 ky , and a subsequen dec ease un il p esen . In
gene al, o he o es species show e y low occu ences du ing his zone,
excep o he las cen u ies, when se e al inc eases in Pinus, Olea and
Que cus a e obse ed. The he baceous componen including A emisia,
As e aceae Cicho ioideae, Poaceae, Plan ago, Ca yophyllaceae and Cis-
aceae, show inc eases du ing his pe iod. The aqua ics and cha coal
concen a ion dec ease conside ably du ing he LH. Wi h espec o he
algae, Bo yococcus dec ease signi ican ly. O he NPP such as Neo-
habdocoela o Glomus inc ease in he las cen u ies.
5. Discussion
Pollen analyses om he Sie a Ne ada ha e been p e iously e-
po ed as p oxies o egional ege a ion and clima e changes (Ande son
e al., 2011; Jim´
enez-Mo eno and Ande son, 2012; Ramos-Rom´
an e al.,
2016, 2018a, 2018b; Mesa-Fe n´
andez e al., 2018; Camue a e al., 2019,
2021). Inc eases in subalpine and empe a e Medi e anean o es spe-
cies in he alpine eco ds, summa ized he e by he PCI (Figs. 5 and 6),
poin o inc eases in humidi y and empe a u e. T ees need ela i ely
high soil wa e supply, while moun ain o es axa mo e o highe
ele a ion du ing clima e wa mings (Lenoi e al., 2008; Go ied e al.,
Fig. 4. De ailed non-pollen palynomo ph (NPP) diag am om he LS-01 co e. The NPP zona ion on he igh was made using clus e analysis p o ided by CONISS
(G imm, 1987). Ligh shadings indica e a 5×exagge a ion scale. HS1, B-A, YD, EH, MH and LH s and o Hein ich S adial-1, Bolling-Alle od, Younge D yas, Ea ly
Holocene, Middle Holocene and La e Holocene.
G. Jim´
enez-Mo eno e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
7
Fig. 5. Compa ison o paleoen i onmen al and paleoclima ic p oxies om he LS-01 co e. (A) P. syl es is ype pollen abundance. (B) Que cus o al pollen in pe -
cen ages. (C) Pollen Clima e Index (PCI; see ex o explana ion). (D) Mic ocha coal concen a ion. (E) C/N a ios om L´
opez-A il´
es e al. (2022). (F) To al O ganic
Ca bon (TOC) om L´
opez-A il´
es e al. (2022). (G) Magne ic suscep ibili y (MS; SI x 10
−4
) om L´
opez-A il´
es e al. (2022). No e he scale is in e ed. (H) Summe
insola ion o 37◦N (Laska e al., 2004). Pollen zones a e shown a he op (see Fig. 3). HS1, B-A, YD, EH, MH and LH s and o Hein ich S adial-1, Bølling-Alle ød,
Younge D yas, Ea ly Holocene, Middle Holocene and La e Holocene.
G. Jim´
enez-Mo eno e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
8
(cap ion on nex page)
G. Jim´
enez-Mo eno e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 631 (2023) 111834
9
Fig. 6. Compa ison o paleoen i onmen al and paleoclima ic p oxies om he LS-01 co e wi h o he paleoclima ic eco ds om he sou he n Ibe ian Peninsula and
summe insola ion o he las 16 ky . (A) Pollen Clima e Index (PCI; see ex o explana ion) om he LS-01 co e. (B) Pedias um and Bo yococcus algae abundances
om LS-01 co e and indica ions abou lake le el and p oduc i i y. (C) Mean Annual P ecipi a ion (MAP) es ima ed om he Padul-15-05 pollen eco d (Camue a
e al., 2022). (D) Pollen Clima e Index (PCI) om he Padul-15-05 pollen eco d (Camue a e al., 2019). (E-G) Sea Su ace Tempe a u e (SST) econs uc ed om
G. bulloides Mg/Ca a ios om ma ine co e ALB-2 (Albo an Sea; Ca al`
a e al., 2019), SST (◦C) econs uc ed om G. bulloides Mg/Ca a ios om ma ine co e
MD95–2043 (Ca al`
a e al., 2019) and SST (◦C) econs uc ed om alkenones om ma ine sedimen co e 293G (Rod igo-G´
amiz e al., 2014). (H) Summe (Augus )
insola ion alues a 37◦N (Laska e al., 2004). Pollen zones a e shown a he op (see Fig. 3). HS1, B-A, YD, EH, MH and LH s and o Hein ich S adial-1, Bølling-
Alle ød, Younge D yas, Ea ly Holocene, Middle Holocene and La e Holocene.
Fig. 7. Compa ison o Pinus pollen eco ds om he Sie a Ne ada wi h Ced us syn he ic eco d om Mo occo o he las 16 ky . (A) Pinus o al pollen om Padul-
15-05 co e (Ramos-Rom´
an e al., 2018a, 2018b; Camue a e al., 2019). (B) Pinus o al and P. syl es is ype pollen abundances om LS-01 co e. (C) Syn he ic Mo occo
z-sco es Holocene Ced us s ack (Jim´
enez-Mo eno e al., 2020). (D) Pinus o al pollen om Laguna de Río Seco, Sie a Ne ada (Ande son e al., 2011). (E-F) Summe
( ed line) and win e (blue line) insola ion alues a 37
◦N (Laska e al., 2004). Pollen zones a e shown a he op (see Fig. 3). HS1, B-A, YD, EH, MH and LH s and o
Hein ich S adial-1, Bølling-Alle ød, Younge D yas, Ea ly Holocene, Middle Holocene and La e Holocene. (Fo in e p e a ion o he e e ences o colou in his igu e
legend, he eade is e e ed o he web e sion o his a icle.)
G. Jim´
enez-Mo eno e al.